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      Middle Pleistocene Hominin Teeth from Longtan Cave, Hexian, China

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          Abstract

          Excavations at the Longtan Cave, Hexian, Anhui Province of Eastern China, have yielded several hominin fossils including crania, mandibular fragments, and teeth currently dated to 412±25 ka. While previous studies have focused on the cranial remains, there are no detailed analyses of the dental evidence. In this study, we provide metric and morphological descriptions and comparisons of ten teeth recovered from Hexian, including microcomputed tomography analyses. Our results indicate that the Hexian teeth are metrically and morphologically primitive and overlap with H. ergaster and East Asian Early and mid-Middle Pleistocene hominins in their large dimensions and occlusal complexities. However, the Hexian teeth differ from H. ergaster in features such as conspicuous vertical grooves on the labial/buccal surfaces of the central incisor and the upper premolar, the crown outline shapes of upper and lower molars and the numbers, shapes, and divergences of the roots. Despite their close geological ages, the Hexian teeth are also more primitive than Zhoukoudian specimens, and resemble Sangiran Early Pleistocene teeth. In addition, no typical Neanderthal features have been identified in the Hexian sample. Our study highlights the metrical and morphological primitive status of the Hexian sample in comparison to contemporaneous or even earlier populations of Asia. Based on this finding, we suggest that the primitive-derived gradients of the Asian hominins cannot be satisfactorily fitted along a chronological sequence, suggesting complex evolutionary scenarios with the coexistence and/or survival of different lineages in Eurasia. Hexian could represent the persistence in time of a H. erectus group that would have retained primitive features that were lost in other Asian populations such as Zhoukoudian or Panxian Dadong. Our study expands the metrical and morphological variations known for the East Asian hominins before the mid-Middle Pleistocene and warns about the possibility that the Asian hominin variability may have been taxonomically oversimplified.

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          Most cited references40

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          A hominid from the lower Pleistocene of Atapuerca, Spain: possible ancestor to Neandertals and modern humans.

          Human fossil remains recovered from the TD6 level (Aurora stratum) of the lower Pleistocene cave site of Gran Dolina, Sierra de Atapuerca, Spain, exhibit a unique combination of cranial, mandibular, and dental traits and are suggested as a new species of Homo-H. antecessor sp. nov. The fully modern midfacial morphology of the fossils antedates other evidence of this feature by about 650, 000 years. The midfacial and subnasal morphology of modern humans may be a retention of a juvenile pattern that was not yet present in H. ergaster. Homo antecessor may represent the last common ancestor for Neandertals and modern humans.
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            A morphometric analysis of maxillary molar crowns of Middle-Late Pleistocene hominins.

            S Bailey (2004)
            This study explores the significance of shape differences in the maxillary first molar crowns of Neandertals and anatomically modern humans. It uses morphometric analysis to quantify these differences and to investigate how the orientation of major cusps, relative cusp base areas and occlusal polygon area influence crown shape. The aims of this study were to 1) quantify these data to test whether the tooth shapes of Neandertals and anatomically modern humans differ significantly and 2) to explore if either of the shapes is derived relative to earlier fossil hominins. Data were collected from digital occlusal photographs using image-processing software. Cusp angles, relative cusp base areas and occlusal polygon areas were measured on Neandertals (n=15), contemporary modern humans (n=62), Upper Paleolithic humans (n=6), early anatomically modern humans (n=3) and Homo erectus (n=3). Univariate and multivariate statistical tests were used to evaluate the differences between contemporary modern humans and Neandertals, while the much sparser data sets from the other fossil samples were included primarily for comparison. Statistically significant differences reflecting overall crown shape and internal placement of the crown apices were found. Neandertals are distinguished from contemporary humans by possessing maxillary first molars that 1) are markedly skewed; 2) possess a narrower distal segment of the occlusal polygon compared to the mesial segment; 3) possess a significantly smaller metacone and a significantly larger hypocone; and 4) possess a significantly smaller relative occlusal polygon area reflecting internally placed cusps. Differences in relative cusp base areas of the hypocone and metacone may contribute to the shape differences observed in Neandertals. However, early anatomically modern humans possessing a pattern of relative cusp base areas similar to Neandertals lack their unusual shape. That the morphology observed in non-Neandertal fossil hominins is more anatomically modern human-like than Neandertal-like, suggests that this distinctive morphology may be derived in Neandertals.
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              Remains of Homo erectus from Bouri, Middle Awash, Ethiopia.

              The genesis, evolution and fate of Homo erectus have been explored palaeontologically since the taxon's recognition in the late nineteenth century. Current debate is focused on whether early representatives from Kenya and Georgia should be classified as a separate ancestral species ('H. ergaster'), and whether H. erectus was an exclusively Asian species lineage that went extinct. Lack of resolution of these issues has obscured the place of H. erectus in human evolution. A hominid calvaria and postcranial remains recently recovered from the Dakanihylo Member of the Bouri Formation, Middle Awash, Ethiopia, bear directly on these issues. These approximately 1.0-million-year (Myr)-old Pleistocene sediments contain abundant early Acheulean stone tools and a diverse vertebrate fauna that indicates a predominantly savannah environment. Here we report that the 'Daka' calvaria's metric and morphological attributes centre it firmly within H. erectus. Daka's resemblance to Asian counterparts indicates that the early African and Eurasian fossil hominids represent demes of a widespread palaeospecies. Daka's anatomical intermediacy between earlier and later African fossils provides evidence of evolutionary change. Its temporal and geographic position indicates that African H. erectus was the ancestor of Homo sapiens.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2014
                31 December 2014
                : 9
                : 12
                : e114265
                Affiliations
                [1 ]Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
                [2 ]National Research Center on Human Evolution (CENIEH), Burgos, Spain
                [3 ]Hexian Museum of Anhui Province, Hexian, China
                [4 ]Anhui Museum, Hefei, China
                [5 ]Chongqing Three Gorges Institute of Paleoanthropology, China Three Gorges Museum, Chongqing, China
                University of Delaware, United States of America
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: MMT WL. Performed the experiments: SX MMT WL. Analyzed the data: SX MMT JMBC YQZ XXF LTZ WBH. Contributed reagents/materials/analysis tools: SX MMT. Contributed to the writing of the manuscript: SX MMT WL.

                Article
                PONE-D-14-18774
                10.1371/journal.pone.0114265
                4281145
                25551383
                32cdae2e-9267-409d-b56a-65e3709ffdf5
                Copyright @ 2014

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 26 April 2014
                : 4 November 2014
                Page count
                Pages: 38
                Funding
                This work has been supported by the grants from Chinese Academy of Sciences (GJHZ201314, KZZD-EW03, and XDA05130100), National Natural Science Foundation of China (41302016 and 41272034), Dirección General de Investigación of the Spanish Ministerio de Educación y Ciencia (CGL2012-38434-C03-02 and Acción Integrada España Francia HF2007-0115), Consejería de Educación de Junta de Castilla y León (CEN074A12-2), and Leakey Foundation. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Paleontology
                Fossils
                Paleoanthropology
                Earth Sciences
                Custom metadata
                The authors confirm that all data underlying the findings are fully available without restriction. All the relevant data that is needed to replicate this study and to draw the conclusions for this study is within the paper and the supporting information files. Data underlying the findings described in the paper (morphological and metric features as well as micro-CT 3D reconstructions) are already provided in the main text and supporting information. The variables that can be extracted from the original micro-CT files regarding the EDJ surface, pulp cavity and tissue proportions are already included in the paper. However, if secondary data or images are needed, or a personal observation on the original fossils is necessary, please contact Dr. Liu ( liuwu@ 123456ivpp.ac.cn ) or Dr. Xing ( xingsong@ 123456ivpp.ac.cn ).

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